The host range and selectivity of a parasitic plant: Rhinanthus minor L.

Summary Rhinanthus minor (Yellow-rattle) is a widespread hemiparasitic plant of grassland habitats throughout Britain. Association analysis of the dune vegetation at Holme-next-the-Sea in eastern England revealed only two potential host plants through positive association. In contrast direct examination of the root systems revealed haustorial connections with 20 host species. The number of species parasitized by one plant ranged from one to seven. Data from another four sites in Britain and one in central Europe indicate that the natural host range of R. minor encompasses at least 50 species from 18 families with 22% in the Leguminosae and 30% in the Gramineae. Comparison of the number of haustorial connections made to each species with the abundance of roots in the soil shows that R. minor is a highly selective parasite, but that the selectivity is not consistent between populations or between plants from different parts of the same population. The reasons for host selectivity are discussed.     

Interactive effects of mycorrhizae and a root hemiparasite on plant community productivity and diversity

Plant communities can be affected both by arbuscular mycorrhizal fungi (AMF) and hemiparasitic plants. However, little is known about the interactive effects of these two biotic factors on the productivity and diversity of plant communities. To address this question, we set up a greenhouse study in which different AMF inocula and a hemiparasitic plant (Rhinanthus minor) were added to experimental grassland communities in a fully factorial design. In addition, single plants of each species in the grassland community were grown with the same treatments to distinguish direct AMF effects from indirect effects via plant competition. We found that AMF changed plant community structure by influencing the plant species differently. At the community level, AMF decreased the productivity by 15–24%, depending on the particular AMF treatment, mainly because two dominant species, Holcus lanatus and Plantago lanceolata, showed a negative mycorrhizal dependency. Concomitantly, plant diversity increased due to AMF inoculation and was highest in the treatment with a combination of two commercial AM strains. AMF had a positive effect on growth of the hemiparasite, and thereby induced a negative impact of the hemiparasite on host plant biomass which was not found in non-inoculated communities. However, the hemiparasite did not increase plant diversity. Our results highlight the importance of interactions with soil microbes for plant community structure and that these indirect effects can vary among AMF treatments. We conclude that mutualistic interactions with AMF, but not antagonistic interactions with a root hemiparasite, promote plant diversity in this grassland community. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Host selectivity and the mediation of competition by the root hemiparasite Rhinanthus minor

Summary Rhinanthus minor (Yellow-rattle) was grown in replacement series mixtures with Lolium perenne and Trifolium repens. The hemiparasitic interaction resulted in Relative Yield Totals (the sum of the yields in mixture relative to those in monoculture) considerably above 2. The hemiparasite caused a greater decrease in the yield of the legume and also performed better on the legume, indicating that T. repens was a better host for R. minor than L. perenne under the experimental conditions. When L. perenne and T. repens were grown in binary mixture with or without R. minor the hemiparasite affected considerably the competitive relationship between the two species by selectively parasitizing the legume. The effect of R. minor on competition between the two species was, however, dependent upon the nutrient status of the soil: the higher the level of soil nitrogen the fewer haustorial connections were made with T. repens and the less was the depression in its yield. In another series of experiments in which Festuca rubra, Holcus lanatus and L. perenne were grown in various binary mixtures with or without R. minor it was also shown that the yield of a preferred host was depressed to the advantage of a non-preferred host. It is suggested that the mediation of competition by the hemiparasite provides a mechanism by which it might affect the structure and diversity of plant communities.     

Seeds of change: The value of using Rhinanthus minor in grassland restoration

Question: What is the value of using Rhinanthus minor in grassland restoration and can restrictions on its establishment be overcome? Location: England (United Kingdom). Methods: Two experiments were established to determine the efficacy of inoculating R. minor on a suite of four agriculturally improved grasslands and the efficacy of using R. minor in grassland restoration. In Experiment 1, the effect of herbicide gap creation on the establishment and persistence of R. minor in grasslands ranging in productivity was investigated with respect to sward management. In Exp. 2, R. minor was sown at 1000 seeds/m² in conjunction with a standard meadow mix over a randomized plot design into Lolium perenne grassland of moderate productivity. The treatment of scarification was investigated as a treatment to promote R. minor. Results: Gap size had a significant role in the establishment and performance of R. minor, especially the 30 cm diameter gaps (Exp. 1). However, R. minor failed to establish long‐term persistent populations in all of the agriculturally improved grasslands. In Exp. 2, establishment of R. minor was increased by scarification and its presence was associated with a significant increase in Shannon diversity and the number of sown and unsown species. Values of grass above‐ground biomass were significantly lower in plots sown with R. minor, but values of total above‐ground biomass (including R. minor) and forb biomass (not including R. minor) were not affected. Conclusions: The value of introducing R. minor into species‐poor grassland to increase diversity has been demonstrated, but successful establishment was dependent on grassland type. The scope for using R. minor in grassland restoration schemes is therefore conditional, although establishment can be enhanced through disturbance such as sward scarification.     

Rhinanthus minor population genetic structure and subspecies: Potential seed sources of a keystone species in grassland restoration projects

In the last few decades unimproved semi-natural grasslands have been affected by intensification of land use and habitat fragmentation. Because of their biodiversity these species-rich grasslands are of high conservation importance and efforts are under way to restore such habitats. Detailed knowledge of within species diversity will aid deciding on the optimal seed source for such restoration projects, e.g. local genotypes or ecotypes. Rhinanthus minor is a species that is typically found in semi-natural grasslands and is commonly used in grassland restoration projects. This is because R. minor is a hemiparasitic plant that takes minerals and nutrients from its host, which in turn decreases the host's biomass and leads to opportunities for less competitive species in the vegetation. Here, we investigate genetic diversity within and between R. minor populations. This allowed us to test whether the six different subspecies of R. minor that have been described in the UK, based on their morphology, flowering time, and habitat, can be differentiated using molecular markers. We identified moderate levels of genetic differentiation between R. minor populations within the UK. In addition, R. minor individuals from the UK appear to be distinct from R. minor and Rhinanthus angustifolius individuals from other European countries based on microsatellite genotyping and DNA sequencing of cpDNA and rDNA ITS. The molecular markers used in the current study did not separate populations of R. minor based on either their subspecies or habitat. The implication for the use of R. minor in grassland restoration projects seems to be that it is not necessary to use local seeds or seeds from the same subspecies.     

Pollination Ecology and Seed Production of Rhododendron Ponticum in Native and Exotic Habitats

Alien plants may be reproductively limited in exotic habitats because of a lack of mutualistic pollinators. However, if plants are adequately served by generalist pollinators, successful reproduction, naturalisation and expansion into exotic habitats may occur. Rhododendron ponticum is very successful, ecologically damaging invasive plant in Britain and Ireland, but is in decline in its native Iberian habitat. It spreads locally by sending out lateral branches, but for longer distance dispersal it relies on sexually produced seeds. Little is known about R. ponticum's pollination ecology and breeding biology in invaded habitats. We examined the flower-visiting communities and maternal reproductive success of R. ponticum in native populations in southern Spain and in exotic ones in Ireland. R. ponticum in flowers are visited by various generalist (polylectic) pollinator species in both native and exotic habitats. Although different species visited flowers in Ireland and Spain, the flower visitation rate was not significantly different. Insects foraging on R. ponticum in Spain carried less R. ponticum pollen than their Irish counterparts, and carried fewer pollen types. Fruit production per inflorescence varied greatly within all populations but was significantly correlated with visitation at the population level. Nectar was significantly depleted by insects in some exotic populations, suggesting that this invasive species is providing a floral resource for native insects in some parts of Ireland. The generality of the pollination system may be factor contributing to R.ponticum's success in exotic habitats.     

Interactions of the Hemiparasitic Species Rhinanthus minor with its Host Plant Community at Two Nutrient Levels

For root hemiparasites, host plants are both the source of water and nutrients below-ground, but competitors for light above-ground. Hemiparasites can reduce host biomass, and in this way considerably affect the whole plant community. To investigate these effects, we carried out two experiments in an oligotrophic meadow with a native population of Rhinanthus minor. In the first experiment, removal of R. minor was combined with fertilization in a factorial design, and in the second one, we manipulated R. minor density by thinning. The presence of R. minor decreased the biomass of its host community, mostly by suppressing grasses. In this way, the species was able to counterbalance the effect of fertilization, which increased community biomass and in particular that of grasses. Neither the presence of R. minor nor fertilization affected the total number of species or the Shannon-Wiener diversity index (H’) of the host community. However, H’ of grasses was higher and H’ of forbs (non-leguminous dicots) was lower in the presence of R. minor. Reduction of grasses by R. minor favored mainly the dominant forb Plantago lanceolata, which partly acquired the role of a competitive dominant. Effects of R. minor on community diversity seem to be highly dependent on the relative sensitivities of dominant and subordinate species. Fertilization increased the mortality of seedlings, resulting in a lower number of flowering plants. However, surviving individuals on average produced more flowers. Thinning resulted in lower mortality of R. minor plants. This indicates that intraspecific competition in R. minor populations results in negative density dependence.     

南京市加拿大一枝黄花入侵地群落的物种组成与多样性特征研究(附表)

为探究外来植物加拿大一枝黄花(Solidago canadensis)入侵与南京市本土植物多样性的关系,该文采用踏查及样方调查对其入侵地群落的物种组成与多样性进行研究。结果表明:(1)入侵地群落中共有维管植物200种(含种下单元),隶属于62科156属,其中被子植物195种,裸子植物1种,蕨类植物4种; 数量最多的是菊科(Asteraceae)和禾本科(Poaceae)植物,分别有25种和24种; 从生活型来看,草本植物占多数,有133种,占所有种的66.50%; 此外,群落内尚有其他外来植物29种。群落中重要值最大的是加拿大一枝黄花,为40.00%; 其次是救荒野豌豆(Vicia sativa),为7.00%。(2)从植物区系看,非入侵地中植物科的区系分布型共4个,而入侵地植物科的区系类型仅有3个,其中泛热带分布、世界分布和北温带分布为两者均有,东亚和北美间断分布型仅在非入侵地中存在。非入侵地植物科的区系以世界分布型为主,有13科,占该类型群落中所有科的39.39%; 入侵地则以泛热带分布型为主,有16科,占总科数的45.71%。非入侵地中植物属的区系分布型有10个,而入侵地有12个,两者区系成分相近,旧世界热带型和热带亚洲至热带大洋洲分布型仅在入侵地中出现。北温带分布型和世界分布型同为两者中最主要成分。入侵地及非入侵地群落属的区系R/T值分别为0.58和0.38,种系分化度分别为3.29和3.11。(3)重度入侵群落的Margalef指数(E)与非入侵及轻度入侵群落相比,显著降低; 此外,重度入侵群落的Simpson指数(D)、Shannon-Weiner(H'')指数和Pielou指数(J)均显著低于非入侵、轻度入侵、中度入侵群落。(4)不同生境之间加拿大一枝黄花群落的E、D、H''和J均无显著性差异。该研究可为南京地区的加拿大一枝黄花入侵地的治理防控和生态恢复,以及进一步的科学研究提供强有力的理论支撑。     

Cedar Infestation Impacts Avian Communities along the Niobrara River Valley,Nebraska

Habitat modification has the potential to cause changes in structure and composition of bird communities. Our goal was to determine the response of Songbird community composition to eastern red cedar (Juniperus virginiana) removal in The Nature Conservancy's Niobrara Valley Preserve, Nebraska. We used point counts to survey birds in the riparian matrix of grassland and forest habitats. More than 60 species were recorded on surveys during 2004–2005. We also use the program PRESENCE to determine the response of five species to various habitat components, including cedar density: House Wren (Troglodytes aedon), Spotted Towhee (Pipilo maculates), Ovenbird (Seiurus aurocapillus), Red‐eyed Vireo (Vireo olivaceus), and Indigo Bunting (Passerina cyanea). Species richness estimates were highest in open and mixed habitat patches. Local populations of Ovenbirds and Red‐eyed Vireos responded positively to cedar density, whereas House Wren numbers declined as cedar density increased. Cedar abundance explained the most variation in bird community similarity between survey points; species richness increased as cedar density decreased. Habitat structure and composition drove variation in community composition and population abundance at fine, local scales within the Preserve. Fine‐scale management to remove cedar from local areas should increase diversity of avian species by maintaining a matrix of habitat types. Cedar removal at any scale will affect the composition of bird communities, and we encourage a structured approach to management decisions.     

Predictive quality of macroinvertebrate — habitat associations in lower navigation pools of the Mississippi River

Macroinvertebrate community structure was compared between habitat types within a navigation pool and between navigation pools of the Upper Mississippi River. Bottom samples were taken using a grab or Wilding sampler from 40 and 14 stations on Pool 19 and 26, respectively. In both Pools, distinct communities developed based on substrate type or the presence of aquatic macrophytes rather than specific habitats as defined by river morphometry. Areas with sand substrates usually had communities of low density and diversity. Communities of the highest density, including Hexagenia or Musculium or both, occurred in habitats with silt-sand substrates. However, these areas were low in diversity. High diversity was found in both vegetated areas and habitats with coarse substrates, the latter dominated by net-spinning caddisfly larvae. Due to pool age and longitudinal distribution of species, community similarity between Pools 19 and 26 was not significant, p < 0.05, but functional feeding similarities occurred between communities from the same type of substrate.     

Responses of insect herbivores to sharing a host plant with a hemiparasite: impacts on preference and performance differ with feeding guild

1. Root hemiparasites are common components of many ecosystems and can affect both the biomass and the nutritional quality of the plants they infect. The consequences of these modifications for the preference and performance of three herbivore feeding guilds sharing a host with the hemi‐parasite were examined. 2. It was predicted that as the hemiparasite increased in biomass its impact on the host would increase, as would the indirect impacts on the herbivores. It was also predicted that herbivores from different feeding guilds would respond differently to the presence of the hemiparasite, reflecting the extent to which they utilise resources disrupted by the parasite and hence are in competition with it. 3. The preference and performance of phloem‐feeding aphids, xylem‐feeding spittle bugs, and leaf‐feeding grasshoppers were measured on the host grass species, Holcus lanatus L. (Poaceae), with and without attachment from the hemi‐parasite, Rhinanthus minor L. (Orobanchaceae). 4. The effects of R. minor on the host were dependent on the hemiparasite's stage of growth, being most pronounced when it was at peak biomass. At this stage it caused a significant reduction in the biomass, water content, and total nitrogen content of the host plants. 5. Overall, herbivores benefited from, or preferred, shared host plants more than uninfected plants. The aphid benefited from sharing a host with R. minor, showing increased population growth on, and preference for, parasitised plants. The spittle bug also showed a preference for parasitised plants. The grasshopper, Chorthippus brunneus Thunberg (Orthoptera: Acrididae), did not show a preference for, or a performance response to, parasitised hosts, but it consumed significantly more plant material when caged on parasitised plants. 6. These data support the prediction that invertebrate herbivores responded to changes in host plant traits driven by the hemiparasite, and strongly suggest that these indirect interactions could impact on population and community processes within natural communities.     

Habitat structure mediates top–down effects of spiders and ants on herbivores

Differences in structural complexity of habitats have been suggested to modify the extent of top–down forces in terrestrial food webs. In order to test this hypothesis, we manipulated densities of generalist invertebrate predators and the complexity of habitat structure in a two-factorial design. We conducted two field experiments in order to study predation effects of ants and spiders and, in particular, of the wasp spider Argiope bruennichi on herbivorous arthropods such as grasshoppers, plant- and leafhoppers in a grassland. Predator densities were manipulated by removal in habitats of higher and lower structural diversity, and the effects on herbivore densities were assessed by suction sampling. Habitat structure was changed by cutting the vegetation to half its height and removing leaf litter.We found a significant negative effect of this assemblage of generalist predators on plant- and leafhoppers, which were 1.6 times more abundant in predator removal plots. This effect was stronger in low-structured (cut) than in uncut vegetation. Densities of the most abundant planthopper Ribautodelphax pungens (Delphacidae) were 2.2 times higher in predator removal plots. Furthermore, adult plant- and leafhoppers responded more strongly than juveniles and epigeic species more strongly than hypergeic species. The presence of predators had a positive effect on plant- and leafhopper species diversity. In a second field experiment, we tested the exclusive impact of Argiope bruennichi on its prey, and found that its effect was also significant, although weaker than the effect of the predator assemblage. This effect was stronger in grass-dominated vegetation compared to structurally more complex mixed vegetation of grasses and herbs. We conclude that habitat structure and in particular vegetation height and architectural complexity strongly modify the strength of top–down forces and indirectly affect the diversity of herbivorous arthropods.     

A comparison of the diversity and structure of butterfly communities in semi-natural and human-modified grassland habitats at the foot of Mt. Fuji, central Japan

To examine the effects of human land use and disturbance on butterfly communities we compared the diversity and structure of communities in relatively undisturbed, semi-natural grassland habitats and highly disturbed, human-modified ones. Comparisons were based on transect counts conducted at 6 study sites at the foot of Mt. Fuji in the cool temperate zone of central Japan during 1995. Out of the six community parameters used in the analyses, the species richness, species diversities H and 1/, and dominance indices were significantly different between the two habitat types stated above. That is, butterfly communities in semi-natural habitats had higher species richness and diversity, and lower dominance indices than those in human-modified ones. This suggests that heavy land modification and disturbance to semi-natural habitats change greatly its butterfly community structure, which, indeed, leads to decreasing species richness and diversity mainly due to the loss of species that are confined to semi-natural habitats. Through the comparisons of various species' characteristics, it was found that the species confined to semi-natural habitats had lower population abundance, fewer generations per year, more restricted local distributions, and narrower geographic range size in Japan than the other component species. Based on our results, it is critical that the persistence of the species that are limited to semi-natural habitats be ensured in order to maintain high species richness and diversity in grassland butterfly communities. Thus, conservation plans that retain as much semi-natural habitat as possible within the process of human grassland use, development, and modification are needed.     

Tibetan sacred sites conserve old growth trees and cover in the eastern Himalayas

Khawa Karpo, in the eastern Himalayas, is a mountain considered sacred throughout Tibet, and is internationally recognized as a global biodiversity hotspot. Numerous areas within this landscape are considered ‘sacred’ by the indigenous Tibetans of the region, who interact with these sites in ways potentially beneficial to conservation. Our previous remote sensing study indicated that sacred sites are found in habitats with greater species richness, diversity, and endemism than randomly selected non-sacred sites. This study examines the role of sanctity in biodiversity conservation within habitats in the Khawa Karpo region by pairing plots within the same habitats in sacred and non-sacred areas. Understory richness, diversity, cover, and number of useful species are measured; for trees, richness, diversity, cover, and density are measured. Results indicate that within habitats sanctity does not affect understory plant communities; however, within sacred areas trees are larger (p = 0.003) and forests have greater cover (p = 0.003) than non-sacred areas. Our results indicate that, whereas placement of sacred areas and preservation of vegetation cover affects useful plants, biodiversity and endemism, within habitats sacred sites preserve old growth trees and forest structure. In sum, Tibetan sacred sites are ecologically unique and important for conservation on varying scales of landscape, community, and species.     

高寒草甸地区小哺乳动物群落多样性的初步研究

高寒苹甸地区的人工草场、半人工草场、次生杂类草草场和撩荒地中共有4个小哺乳类群落;根田鼠+高原鼢鼠群落;根田鼠+甘肃鼠兔群落;高原鼠兔+高原鼢鼠群落;长尾仓鼠+高原鼠兔群落。小哺乳类群落种的多样性与植物群落种的多样性无显著相关关系,而与植被的高度、盖度呈显著负相关关系。高原鼢鼠是各生境中的广布种。根田鼠与甘肃鼠兔在植被郁闭度高的环境中为群落的主要组成者;植被郁闭度低的环境中,高原鼠兔和长尾仓鼠是群落的主要成员。它们的空间格局主要反映空间环境条件的差异。     

The Arrhenatheretum on set-aside at Aldroughty,Elgin

Summary

The Arrhenatheretum on set-aside in two fields at Aldroughty Farm, Elgin, is described. This vegetation is noteworthy because some of the attractive grassland and wetland species in it are very rare on Scottish set-aside. The species involved (e.g. Carex flacca, Dactylorhiza purpurella, Rhinanthus minor) are generally entirely absent from lowland farmed landscapes, but populations do occur in areas of semi-natural habitat at Aldroughty. It is concluded that, as far as most long-term set-aside is concerned, vegetation of greater conservation value does not develop thereon owing to non-availability of the necessary seeds.     

垫状植物囊种草对群落物种多样性的影响

在甘肃盐池湾国家级自然保护区内海拔4137 m处,选择典型的囊种草垫状植被设置研究样地,研究了垫状植物囊种草对群落物种组成和群落物种多样性的影响,并且定量的研究了囊种草对群落物种丰富度的影响能力和维持潜力。研究结果表明:囊种草为群落中增加了新的植物种类,并且提高了部分生境一般种的多度;囊种草的出现提高了群落物种密度和物种丰富度,进而提高了群落物种多样性;囊种草斑块的增加将会引起景观水平物种丰富度的增加,表明囊种草具有为群落中引入新的植物种类进而提高群落物种丰富度的能力;在景观水平,囊种草所创造的生境多样性则成为一种保障,可以维持景观中物种丰富度从而降低物种损失的风险,表明囊种草具有较高的群落物种丰富度维持潜力。     

Grassland plant species occurring in extensively managed road verges are filtered by urban environments

Background: Urbanisation filters species in communities depending on their adaptability to conditions in built-up areas, especially in semi-natural habitats. Roadside vegetation is widespread along urban-rural gradients and is therefore a good place to study landscape-scale factors influencing plant community composition.

Aim: Our study aimed to assess how plant species distributions vary between urban and rural landscape contexts and to identify biological traits favoured in urban areas.

Methods: Presence/absence data for 63 indigenous common species were collected in 296 road verge patches distributed along the urban–rural gradient in three French cities. We investigated the effects of landscape composition on species assemblages and related individual species responses to urbanisation to functional traits associated with dispersal and persistence capacity.

Results: Many grassland species were negatively affected by increasing proportion of built-up areas in the landscape. Insect-pollination and high seed production appeared to be key traits favoured in grassland communities in urban areas, whereas dispersal modes were less related to plant distribution.

Conclusions: This study has demonstrated that urban filters affect common species of widespread, managed road verges. Better knowledge of the flora of these herbaceous roadsides may contribute to the conservation of common biodiversity within other grassland habitats found in urban areas.     

Habitat associations of epigeal spiders in upland calcareous grassland landscapes: the importance for conservation

Upland calcareous grassland landscapes are typically comprised of a matrix of calcareous grassland, acid grassland and limestone heath plant communities. This matrix of habitats is produced by a combination of underlying geology, climate and management. These landscapes are typically managed through grazing, with management targeted to maintain particular plant communities in the calcareous grassland habitat, whilst patches of acid grassland and limestone heath are not targeted by conservation management. The biodiversity value of acid grassland and limestone heath patches within the calcareous grassland matrix are unknown. This study provides the first assessment of their biodiversity value by examining aspects of epigeal spider diversity supported by these non-target habitat patches in comparison to calcareous grassland. Spiders were sampled in each habitat from April to August 2014 using pitfall traps across three upland regions in Great Britain. Spider species assemblages were distinct between limestone heath and both grassland types. Distinction in species assemblages are likely due to differences in vegetation structure and microclimate, e.g., humidity, degree of shade. Each habitat type supported several rare species (e.g., Jacksonella falconeri, Agyneta subtilis) revealing the contribution to spider fauna. The distinct spider species assemblage and presence of rare species in limestone heath patches demonstrate their importance in the upland calcareous grassland matrix. This study highlights the value of monitoring biodiversity in non-target habitats within a habitat matrix alongside those that are actively targeted by management.     

Pattern of land mosaics affecting butterfly assemblage at Mt Ikoma, Osaka, Japan

We studied the effects of habitat mosaics on butterfly assemblage on multiple spatial scales: landscape, landscape element, local habitat, and microhabitat, based on the transect counts conducted along a 3.84 km route. The transect route, including 21 local habitats, passed through two distinct areas: 1.65 km of a secondary deciduous Quercus forest and the grove of a shrine in Hiraoka, and 2.19 km of a mosaic of secondary deciduous Quercus forest, grassland, and farmland in Narukawa. The diversity of the landscape elements and species richness were higher in Narukawa than in Hiraoka; the landscape mosaic enhanced the species richness in Narukawa. However, the diversity indices and specialist species (univoltine tree feeder) were decreased in this mosaic landscape. The species richness at local habitats was also increased by the mosaic of microhabitats, such as the herbaceous layer, glade, and mantle in the local habitats, whereas it was decreased by an abundant shrub layer. The ratios of species richness to abundance in the local habitats were lower than expected based on random sampling from the total of Hiraoka and Narukawa. This means that local assemblages were non-random samples from an assemblage on the landscape or regional scale, and were made up by the process of habitat selection of butterfly species in the assemblages on the landscape or regional scale. For conservation of butterfly assemblages, we recommend that woodlands should be kept without fragmentation, but with glades or small grasslands, and with clearance of the shrub layer along the path.